Unit 11:  Sub-Phylum Vertebrata,  The Fish

1.  Briefly describe the excretory, skeletal, muscle and nervous systems and method of feeding of the supposed immediate ancestor of the first vertebrate, and relate each of these characteristics to its function in the evolution of fish.
2.  Relate fish evolution to the geologic time in which it occurred, including the names and appearance of the oldest fossil fish.
3.  Create a fish family tree showing the relationship of the Ostracoderms, Cyclostomata, Placoderms, Chondrichthyes and Osteichthyes.
4.  Similarly to objective #4, draw a family tree of the Osteichthyes showing the relationship between the lobefins and the ray fins;  describe any outstanding features of each group.
5.  Describe the basic differences between pronephros, mesonephros and metanephros kidneys, and relate eaach type to the vertebrate group in which it is found.
6.  Describe the external appearance of lampreys and hagfish.
7.  Outline the life cycle of the sea lamprey, including the apparent trigger for metamorphosis.
8.  List the ways in which the Chondrichthyes are more advanced than the cyclostomes.
9.  Describe the heterocercal tail of the shark.  Describe in detail how the body plan of the shark helps it achieve lift.
10.  Describe the respiratory apparatus of the shark and special problems it encounters because it lacks an operculum.
11.  Compare the most common reproductive patterns in sharks and in bony fish.
12.  State the original function of the swim bladder, and describe the extent to which it is still used for that function in the different osteichthyes.
13.  Describe the anatomy and function of the lateral line system, and name the fish groups which have this organ.
14.  Compare and contrast the homocercal tail of the bony fish with the heterocercal tail of the sharks.
15.  Describe the differences between ram ventilation, the usage of spiracles and the countercurrent mechanism employed by fish to accomplish respiration.
16.  Name the fish groups in which the notochord is completely replaced by vertebrae.
17.  Name and describe the characteristics of the two main body patterns in the teleosts, and give examples of local fish representing each body pattern.
18.  List and briefly describe the ways in which the bony fish are more advanced than the shark group and cyclostomes.
19.  Define catadromous and anadromous and give examples organisms described by these terms.

Resources:  Text:  Animal Diversity, Chapter 16
Internet Resources:
    http://www.ucmp.berkeley.edu/vertebrates/basalfish/chondrintro.html
    http://www.biology.ualberta.ca/jackson.hp/iwr/taxa/Chondrichthyes.html
    http://www.seaworld.org/Fishes/class.html
    http://www.mhhe.com/hickmanad4e

FISH:  CLASS CHONDRICHTHYES,  CLASS OSTEICHTHYES
    -16th century included seals, whales, amphibians, crocodiles and hippos;
        jellyfish, starfish etc

    **Today:  gill breathing, ectothermic, aquatic vertebrates that posses fins and
        have skin   usually covered w/scales; masters of aquatic environment
    -humans likely to be uninformed about fishes:

        1. environment alien to humans
        2. most nature shows and books devote time and space to birds/mammals

    -~23,000 species, more than all other vertebrates put together

    Evolution- may have arisen from free swimming larval tunicate
        Neotony- ability of a larval form to reproduce, larval tadpole form will escape adult
            sessile form

    CLASSIFICATION- ** not even a majority of ichtyologists agree on plan

            Subphylum Vertebrata
                    Superclass Agnatha (no jaw)- hagfish/lamprey

                    Superclass Gnathostomata (jawed fishes)

                            Class Chondrichthyes- skeleton of cartilage
                                -no operculum (gill cover); no swim bladder
                                -sharks, skates, rays

                            Class Osteichthyes- bony skeleton (teleostomi)
                                -have operculum, swim bladder

                                    Subclass Actinopterygii- ray finned fish
 

CHARACTERISTICS:
Superclass Agnatha
    -eel-like
    -no paired appendages
    -notochord persists
    -biting mouth; predatory/parasitic existance
    -cartilagenous cranium
    -simple heart; one atrium, one auricle, aortic arches in gill region
    -dorsal nerve cord
    -16 pairs of gills in hagfish; 7 pr in lampreys

    -digestive system w/o stomach; spiral valve separates posterior end of esophagus
        from  straight intestine; typhlosole partitions intestine in spiral manner to increase
        surface area

    -fertilization external; hagfish hermaphroditic (rare in vertebrates) but only one gonad
        functional; lampreys dioecious w/larva and metamorphosis

    -kidneys- in vertebrates differ in relationship to blood system, complexity and efficiency
        (maintain same mineral levels inside as those present outside)

        Pronephric- most primitive, in embryonic stages of all vertebrates, functional in adults
            of none
 
 
 
 
 
 

       Mesonephric- fish and amphibians
 
 
 
 
 

       Metanephric- reptiles, birds, mammals
 
 
 
 
 
 

Lamprey life cycle
    -feed as adults in open ocean, near end of life migrate to freshwater streams to mate

    -male begins nest building and then joined by female
    -female attaches to stone w/mouth to maintain position over nest, male attaches to
        dorsal side of female head
    -egg and sperm shed into nest
    -eggs stick to pebbles and become covered by sand

    -eggs hatch into ammocoetes larva; ~2 weeks burrow into sand or mud; live as filter
        feeders

    -metamorphosis can last 3-7 years!!  at time of metamorphosis endostyle becomes a
         thyroid gland and chemical stimulation   by thyroxin triggers metamorphosis

Class Chondrichthyes
    -sharks, skates, rays ~760 species
    -predaceous/carnivorous, scavengers

    -biting mouth parts located ventral, paired appendages
    -cartilaginous skeleton

    -no swim bladder; sharks heavier than water, therefore, must remain in motion to avoid
        sinking
            Heterocercal tail w/vertebrae in dorsal lobe gives lift; streamlining w/flat ventral
                surface can also aid lift
 
 
 
 
 
 

    -no operculum (gill cover), 7 pr exposed gills

    -tough skin w/ placoid scales.  Toothlike structures w/ dentin coverings; scales project
        toward posterior to give sandpaper texture (posterior projections reduces friction)

    -teeth modified placoid scales.  Row of teeth on outer edge of jaw backed up by rows
        of teeth  attached to a ligamentous band.  As outside teeth wear down or break they
        are replaced by  newer teeth

    -dioecious; internal fertilization; oviparous, ovoviviparous and viviparous (?)

    -secondary facilities: smell, lateral line system; bioelectric sense

    -largest fish- whale shark, longer than 10m, not predatory; filter feeder
    -most feared; great white (Jaws)
        27 of 760 species known to attack man (often terretorial confrontation or provocation)

Class Osteichthyes- bony fish
    -skeleton (at least partly) made of ossified bone

    -swim bladder for bouyancy; gill w/ operculum
    -body form great example of adaptive radiation
            (*one form generating many variations dependent upon niche)

    -skin w/ scales:     ganoid- tile like, diamond shape, little overlap
                                 cycloid - smooth, light, thin, flexible, overlap
                                 ctenoid - rough, light, thin, flexible, overlap

                                   or none ---> catfish

    -terminal mouth
    -dioecious, external fertilization; oviparous

    -Super order Teleostei ~ 21,800 species, represent 96% of all fish
    -homocercal tail- upper and lower lobes nearly equal in size
 
 
 
 
 
 

    -lungs of primitive groups (lung fish) evolved into swim bladder
    -scale growth discloses seasonal changes in growth rate, much like trees; scale growth
        proportional to increase in length

GENERAL ADVANCES:
1. locomotion
    -design to overcome resistance encountered in dense water

    -less energy expenditure for a fish to swim than a terrestrial animal to run
    -friction reduced between fish and water by streamlined shape and mucus secretions
        that  lubricate body surface

    -bouyancy properties of water also contribute to efficiency; fish spend little energy
        combatting  the pull of gravity
    -fishes move using fins and body wall to push against water

    -body wall muscles arranged in bundles called myotomes

    -anyone who has eaten fish recognize muscle layers that are arranged in a
 
 
 

                muscles extend posteriorly and anteriorly in zig zag fashion, contraction of
                    myotome effects large portion of body wall.

    -myotomes on opposite sides of body contract alternately; vertebral column prevents
        body from collapsing; body flexes laterally.  Waves of contraction pass along body
        from anterior ---> posterior
 
 
 
 

    *lampreys, hagfish and eels swim inefficiently at best; use of fin minimal
    *tuna, mackral- very efficient, fast; supplement w/ caudal fin that is tall and forked
        (forked  shape reduces surface area that could cause turbulance)

    *backward swimming possible using pectoral and pelvic fins and reversal of muscular
        waves,  little used and poorly executed

GAS EXCHANGE
    -environment of fish contains 2.5% of the oxygen in air, therefore, large quantities of water
        must pass over gill

    -muscles around pharynx and opercular cavity power pump that moves water into mouth
            and  over gill
            *operculum closed/ mouth open ---> water drawn into pharynx, hydrostatic pressure
                draws water across gills

            *mouth closed/ operculum opens ---> water out

    -Sharks, open ocean bony fish maintain water flow by holding mouth open while swimming,
        RAM VENTILATION

    -spiracles (sharks) are modified gill slits behind eye; used as an alternate route for water
        entering pharynx

    -Exchange very efficient; exchange at gill lamellae
          Counter current mechanism- water and blood move in opposite directions on either
            side of lamellae

            -ensures the amount of oxygen is water always greater than oxygen  in blood,
                therefore, always maintains diffusion gradient

              BLOOD OXYGEN %
 
 
 
 

                                                                                                                      WATER OXYGEN %
 

                Parallel system

              BLOOD OXYGEN %
 
 
 

                                                                                                                       WATER OXYGEN %

        Bouyancy- fish maintain verticle position in water in 4 ways:

            1.  incorporate low density compounds into tissues; fish (especially livers) are
                    saturated    w/ bouyant oils.

            2. use fins to provide lift (caudal fin of shark)

            3. reduce heavy tissue- shark cartilage; fish bone less dense than terrestrial animals

            4. swim bladder- regulate volume of gas; gas secreted into bladder from blood;
                    some    get air directly into bladder through pneumatic ducts from esophagus.
 

NERVOUS AND SENSORY FUNCTIONS

    -CNS of brain and spinal cord, sensory receptors all over body
        Receptors for touch, temp., vision, olfaction, balance, water movement

        Salmon/lampreys rely heavily on smell to return home

    -eyes similar to other vertebrates, most lidless, focus by moving lens up or back in eye
        (most other vertebrates change shape of lens)

    -lateral line system- sensory pits in epidermis of skin connecting canals running
            below epidermis; receptors  stimulated by water moving against them; used to
            detect prey or predator; low frequency sound may also be detected (vibrations)

    -Electroreception- detection of electric fields;  demonstrated in over 500 species of
            fish in   seven families of Chondrichthyes and osteichthyes

        *used to detect prey or to orient toward or away from objects; best known in sharks
            and   rays
 

REPRODUCTION AND DEVELOPMENT
    -millions of eggs produced in a single season
    -numbers reflect hazards of developing in aquatic habitats unattended by parent:
        Vast majority will not survive to reproduce
            -many eggs never get fertilized

            -fertilized eggs may wash ashore and dry

            -many eggs and embryo may be smashed by currents or tides

            -predation

            *if only four of the millions survive the pop. will double

     -large #s not only strategy:  mating behavior, nesting behavior, help protect against
        predation, sedimentation, fouling

    -mating can occur in schools (spawning pheromone)

    -vast majority of fish are oviparous (w/ external fertilization) Exceptions: guppies,
        some sharks ovoviviparous

            grey sharks, hammerheads are viviparous (modified oviducts diverts nutrients
                from female to yolk sac of developing embryo)

                    *sharks fertilize internally

    -male sharks have modified pelvic fin, clasper; during copulation a clasper is inserted
        into  cloaca of female;  sperm travel along groove on clasper.  Result in higher
        proportion of eggs  being fertilized, therefore, fewer eggs

    -few cases of shortterm parental care after birth but for most part they are on their own
            Sticklebacks; sunfish

    -growth temperature dependent
            Temperate regions ---> rapid growth in summer, little in winter reflected in scales

    -migratory fish
           Catadromous- most of life in FW, spawn in salt water (eel)

           Anadromous- most of life in salt water, spawn in FW (salmon)

            *guided by odors, imprinted when migration to sea begins.  Create maps of odors as
                guide upon return

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